Torque vectors of neck muscles in the cat

Summary Anatomical texts describe the neck musculature without measurements of muscle locations or quantitative estimates of pulling actions (torques). This study is based on measurements in stereotaxic coordinates of cat neck muscle origins and insertions, and neck intervertebral rotation axes. Torque vectors in three dimensions were calculated for 14 pairs of dorsal and ventral muscles that insert on the skull or first cervical vertebra. Predicted torque vectors were in general agreement with qualitative statements in the literature. Biventer cervicis and the rectus capitis major, medius, and minor muscles act mainly to raise the head, and longus capitis acts almost exclusively to lower the head. Longissimus capitis, sternomastoid, and cleidomastoid act mainly to roll the head. Complexus acts about equally to raise the head and roll it. Splenius and occipitoscapularis have torque in all three coordinate directions. Torques were altered by changing the pitch of the head with respect to the neck. The calculated neck muscle torques did not correspond to previously reported directions of neck muscle excitation during the vestibulocollic reflex. The neck musculature appears to be a complex, multidimensional system that presents interesting problems in motor control.     

Electromyographic studies of neck muscles in the intact cat

Summary Natural head movements in alert, unrestrained cats were studied using video-filming, videofluoroscopy and electromyographic (EMG) recording methods. In each cat, up to sixteen neck muscles or neck-muscle compartments were implanted with recording electrodes. Patterns of muscle recruitment were examined during systematically-selected behavioral epochs in which the cat held a range of stationary postures, and when it performed volitional and exploratory behaviors such as flexion-extension or turning, grooming, eating, or headshaking. Patterns of muscular activity were interpreted with reference to simultaneous video images of head and neck movements. In separate, videofluoroscopic analysis, flexion-extension movements were examined to gain insight into the underlying movements of the skull and cervical vertebrae. These and other movements were found commonly to depend upon changes in joint angles between lower as well as upper cervical joints. Stationary postures in which the neck was held vertically were consistently associated with tonic EMG activity in only two long dorsal muscles, biventer cervicis and occipitoscapularis. Less consistent activity was also present in dorsal intervertebral muscles crossing lower cervical joints. When the neck was held horizontally, the long dorsal muscles increased their EMG activity and moderate activity was also recorded in deeper intervertebral and suboccipital muscles. When flexion-extension occurred around upper cervical joints, greatest activity was recorded in rectus capitis posterior and complexus, but when it involved the lower cervical joints, large changes in EMG activity could also be detected in biventer cervicis, occipitoscapularis, and the intervertebral muscles crossing lower cervical joints. During specialized, sagittal-plane movements such as grooming, well-defined patterns of synergy could be recognized that varied according to the degree of involvement of upper and lower cervical joint-sets. Movements in the horizontal plane were associated with EMG activity in a largely different subset of neck muscles including splenius, longissimus capitis and obliquus capitis inferior. The levels of EMG activity during flexion-extension or turning movements were much lower than those observed during other more vigorous behaviors, such as head shaking. Some neck muscles, such as clavotrapezius and sternomastoideus, could only be recruited during forceful or ballistic head movements. Results showed that the patterns of muscular activation were linked not only to the speed and trajectory of the movements of the skull, but also to the kinematics of the motion occurring across different parts of the cervical column.     

Effect of neck posture on patterns of activation of feline neck muscles during horizontal rotation

The electromyographic (EMG) patterns of neck muscles were recorded during whole-body horizontal rotation in head-free, alert cats and head-restrained, decerebrate cats. In some trials the cervical column of the animal was oriented vertically, whereas in others it was oriented more horizontally. In alert cats making head movements that compensated for the motion of the platform, neck muscles with modulated patterns of activity could be divided into a subset whose individual EMG patterns changed significantly when the neck posture was altered (including longissimus capitis, obliquus capitis superior and scalenus anterior) and a subset whose individual EMG patterns were invariant regardless of neck posture (including obliquus capitis inferior, levator scapulae and complexus). In head-restrained, decerebrate cats, electromyograms from all implanted muscles were modulated similarly in phase with the platform position. Changing the orientation of the neck had little effect upon these EMG patterns evoked by the horizontal vestibulocollic reflex. One decerebrate cat with strong extensor tone was tested further under head-free conditions. There was very little compensatory head movement, but individual neck muscles displayed patterns of activity that were more similar to those observed in alert, head-free animals.     

Dynamic properties of the vertical otolith neck reflexes in the alert cat

Summary Electromyographic activity of dorsal neck muscles elicited by sinusoidal vertical linear accelerations was studied in alert cats over a wide range of frequencies. Experiments were performed in headfixed cats and total darkness in order to activate selectively the otolith system. The polyunitary EMG activity was recorded from splenius capitis muscles in normal and labyrinthectomized cats during vertical translations varying from 0.05–1 Hz with a fixed 290 mm peak-to-peak amplitude. The corresponding accelerations ranged from 0.003–1.2 g. In normal cats, the results showed a bilateral and sinusoidal modulation of the EMG activity characterized by two typical EMG patterns depending on the stimulus frequency. In the low-frequency range (0.05–0.25 Hz), the neck muscles responses were composed of a second harmonic (frequency double that of the input signal: H2 responses). The H2 pattern was characterized by an increase in EMG activity during both the upward and downward parts of translation. These two components of the H2 response were closely related to the two peak velocities (+90° and –90°) of the animal motion. Only slight decreases in amplitude and shifts in phase were observed when increasing the frequency. In the higher frequency range (0.25–1 Hz), the neck muscles response was composed of a fundamental frequency corresponding to the input signal (H1 response). The H1 pattern was in phase with the peak of downward acceleration at 0.25 Hz. A phase lag (up to 45°) and a gain attenuation (16.5 dB) were observed when increasing the frequency. The two H1 and H2 EMG patterns were totally absent in bilateral vestibular neurectomized cats. In unilateral vestibular neurectomized cats, a strong drop in gain and phase advance was noted, which mainly affected the H1 pattern. The present results describe some characteristics of otolith-spinal reflexes acting on the head musculature during vertical motion. They are compared with the neuronal responses that we have recorded within the vestibular nuclei complex in the same experimental conditions. The functional role of the vertical otolithneck reflexes in stabilizing the head in space during many real-life situations is discussed.     

Effect of gaze direction on neck muscle activity during cervical rotation

Control of the neck muscles is coordinated with the sensory organs of vision, hearing and balance. For instance, activity of splenius capitis (SC) is modified with gaze shift. This interaction between eye movement and neck muscle activity is likely to influence the control of neck movement. The aim of this study was to investigate the effect of eye position on neck muscle activity during cervical rotation. In eleven subjects we recorded electromyographic activity (EMG) of muscles that rotate the neck to the right [right obliquus capitis inferior (OI), multifides (MF), and SC, and left sternocleidomastoid (SCM)] with intramuscular or surface electrodes. In sitting, subjects rotated the neck in each direction to specific points in range that were held statically with gaze either fixed to a guide (at three different positions) that moved with the head to maintain a constant intra-orbit eye position or to a panel in front of the subject. Although right SC and left SCM EMG increased with rotation to the right, contrary to anatomical texts, OI EMG increased with both directions and MF EMG did not change from the activity recorded at rest. During neck rotation SCM and MF EMG was less when the eyes were maintained with a constant intra-orbit position that was opposite to the direction of rotation compared to trials in which the eyes were maintained in the same direction as the head movement. The inter-relationship between eye position and neck muscle activity may affect the control of neck posture and movement.     

Neck muscle responses to stimulation of monkey superior colliculus. I. Topography and manipulation of stimulation parameters

The role of the primate superior colliculus (SC) in orienting head movements was studied by recording electromyographic (EMG) activity from multiple neck muscles following electrical stimulation of the SC. Combining SC stimulation with neck EMG recordings provides an objective and sensitive measure of the SC drive onto neck muscle motoneurons, particularly in relation to evoked gaze shifts. In this paper, we address how neck EMG responses to SC stimulation in head-restrained monkeys depend on the rostrocaudal, mediolateral, and dorsoventral location of the stimulating electrode within the SC and vary with manipulations of the eye position prior to stimulation onset and changes in stimulation current and duration. Stimulation predominantly evoked EMG responses on the muscles obliquus capitis inferior, rectus capitis posterior major, and splenius capitis. These responses became larger in magnitude and shorter in onset latency for progressively more caudal stimulation locations, consistent with turning the head. However, evoked responses persisted even for more rostral stimulation locations usually not associated with head movements. Manipulating initial eye position revealed that the magnitude of evoked responses became stronger as the eyes attained positions contralateral to the side of stimulation, consistent with a summation between a generic command evoked by SC stimulation and the influence of eye position on tonic neck EMG. Manipulating stimulation current and duration revealed that the relationship between gaze shifts and evoked EMG responses is not obligatory: short-duration (<20 ms) or low-current stimulation evoked neck EMG responses in the absence of gaze shifts. However, long-duration stimulation (>150 ms) occasionally revealed a transient neck EMG response aligned on the onset of sequential gaze shifts. We conclude that the SC drive to neck muscle motoneurons is far more widespread than traditionally supposed and is relayed through intervening elements which may or may not be activated in association with gaze shifts.     

Short-term changes in neck muscle and eye movement responses following unilateral vestibular neurectomy in the cat

The purpose of this study was to investigate changes in neck muscle and eye movement responses during the early stages of vestibular compensation (first 3 weeks after unilateral vestibular neurectomy, UVN). Electromyographic (EMG) activity from antagonist neck extensor (splenius capitis) and flexor (longus capitis) muscles and eye movements were recorded during sinusoidal visual and/or otolith vertical linear stimulations in the 0.05–1 Hz frequency range (corresponding acceleration range 0.003–1.16 g) in the head-fixed alert cat. Preoperative EMG activity from the splenius and longus capitis muscles showed a pattern of alternate activation of the antagonist neck muscles in all the cats. After UVN, two motor strategies were observed. For three of the seven cats, the temporal activation of the individual neck muscles was the same as that recorded before UVN. For the other four cats, UVN resulted in a pattern of coactivation of the flexor and extensor neck muscles because of a phase change of the splenius capitis. In both subgroups, the response patterns of the antagonist neck muscles were consistent for each cat independently of the experimental conditions, throughout the 3 weeks of testing. Cats displaying alternate activation of antagonist neck muscles showed an enhanced gain of the visually induced neck responses, particularly in the high range of stimulus frequency, and a gain decrease in the otolith-induced neck responses at the lowest frequency (0.25 Hz) only. By contrast, for cats with neck muscle coactivation, the gain of the visually induced neck responses was basically unaffected relative to preoperative values, whereas otolith-induced neck responses were considerably decreased in the whole range of stimulation. As concerns oculomotor responses, results in the two subgroups of cats were similar. The optokinetic responses were not affected by the vestibular lesion. On the contrary, otolith-induced eye responses showed a gain reduction and a phase lead. Deficits and short-term changes after UVN of otolith- and semicircular canal-evoked collic and ocular responses are compared. Received: 15 April 1997 / Accepted: 29 December 1997     

Cross-innervation of fast and slow-twitch muscles by motor axons of the sural nerve in the mouse

The extensor digitorum longus (EDL) or soleus muscles of adult mice were cross-innervated by the sural nerve (SN) and deprived of their original innervation. The number and sizes of motor units and the location of endplates in these muscles were studied 1.5 to 16 months later. In the EDL muscle, the SN cross-innervated the original endplates. Very few ectopic endplates were seen, even when the nerve was implanted well outside of the original endplate area. Only 3% of the fibres were polyneuronally innervated. In the soleus muscle, however, the SN formed large numbers of ectopic endplates whether the nerve was implanted in the original endplate zone or outside of it. In addition, 20% of the muscle fibres were polyneuronally innervated. The SN cross-innervated both EDL and soleus muscles completely. There was no preference for a particular group of the SN motoneurones since all the cross-innervated muscles were innervated by all SN motor axons and the motor unit sizes of the SN were similar in the cross-innervated EDL and soleus muscles. It is concluded that intrinsic properties of a muscle determine the ability to form ectopic synapses. The distribution of the motor unit sizes is determined by the particular pool of motoneurones which innervates the muscle.     

EMG activities of neck muscles underlying lateral flexion of the neck during head-turning induced by electrical stimulation of the caudate nucleus in cats

Patterns of EMG activities of neck muscles underlying the initiation of head-turning, induced by stimulation of the caudate nucleus, were analyzed with special reference to temporal relations between the onset of head-turning and that of changes in EMG activities. These patterns were compared with those associated with the initiation of lateral flexion of the neck which occurred without electrical stimulation of the caudate nucleus in order to examine whether the caudate-induced head-turning was initiated via the same muscular system as that used in non-caudate-induced head movements. Experiments were carried out using 5 awake, unrestrained cats which were trained to stand still with one limb on each of 4 footplates. Trains of stimulating current pulses were applied to several stimulation points in the caudate nucleus while the animal maintained a stable standing posture with its neck extended. Head movements in the horizontal plane and EMGs of 6 neck muscles (splenius, longissimus cervicis, obliquus capitis caudalis, biventer cervicis, complexus and cervical multifidus) were recorded. Patterns of EMG activities of neck muscles around the onset of the caudate-induced head-turning were characterized by an increase in activity of the splenius, the longissimus cervicis and the obliquus capitis caudalis muscles, and by a decrease in activity of the complexus, the biventer cervicis and the cervical multifidus on the side of flexion. It is suggested that an increase in activity of the splenius, the longissimus cervicis and the obliques capitis caudalis muscles was responsible for the initiation of this evoked response. In non-caudate-induced lateral flexion of the neck, patterns of activities of neck muscles were similar to those in caudate-induced head-turning. It is therefore concluded that the caudate-induced head-turning as an evoked behavioral response was initiated through a muscular system similar to that utilized for similar head movements occurring without electrical stimulation of the caudate nucleus, although the pathways involved are thought to be different.     

Viscosity of the elbow flexor muscles during maximal eccentric and concentric actions

The aim of the present study was to estimate the damping coefficient (B factor) of the elbow flexor muscles during both eccentric and concentric muscle actions. We used a muscle model consisting of a viscous damper associated in parallel with a contractile component, both in series with an elastic component. The viscous damper allowed the concentric loss and the eccentric gain of force to be modelled. Eight volunteer subjects performed maximal eccentric and concentric elbow movements on an isokinetic dynamometer at angular velocities of 0.52, 1.04 and 2.09 rad·s^–1. Torques at an elbow joint angle of 90° were recorded. Electromyogram (EMG) signals from the belly of the right elbow flexor and from the long head of the triceps brachia muscles were recorded using two pairs of bipolar surface electrodes. The root mean square (rms) of the EMG was determined. Eccentric and concentric rms were not significantly different (P>0.05). The B factor was higher in the concentric than in the eccentric conditions (P<0.05), and, whatever the muscle action type it decreased as the velocity increased. These results indicated that the concentric loss and the eccentric gain of force were attributable to the behaviour of the contractile machinery. Furthermore, whatever the exact cause of loss and gain of tension, our study showed that the total effect can be modelled by the viscous damper of a three-component muscle model.     

Neural control of superficial and deep neck muscles in humans

Human neck muscles have a complex multi-layered architecture. The role and neural control of these neck muscles were examined in nine seated subjects performing three series of isometric neck muscle contractions: 50-N contractions in eight fixed horizontal directions, 25-N contractions, and 50-N contractions, both with a continuously changing horizontal force direction. Activity in the left sternocleidomastoid, trapezius, levator scapulae, splenius capitis, semispinalis capitis, semispinalis cervicis, and multifidus muscles was measured with wire electrodes inserted at the C(4)/C(5) level under ultrasound guidance. We hypothesized that deep and superficial neck muscles would function as postural and focal muscles, respectively, and would thus be controlled by different neural signals. To test these hypotheses, electromyographic (EMG) tuning curves and correlations in the temporal and frequency domains were computed. Three main results emerged from these analyses: EMG tuning curves from all muscles exhibited well-defined preferred directions of activation for the 50-N isometric forces, larger contractions (25 vs. 50 N) yielded more focused EMG tuning curves, and agonist neck muscles from all layers received a common neural drive in the range of 10-15 Hz. The current results demonstrate that all neck muscles can exhibit phasic activity during isometric neck muscle contractions. Similar oscillations in the EMG of neck muscles from different layers further suggest that neck motoneurons were activated by common neurons. The reticular formation appears a likely generator of the common drive to the neck motoneurons due to its widespread projections to different groups of neck motoneurons.     

Cross-validated models of the relationships between neck muscle electromyography and three-dimensional head kinematics during gaze behavior

The object of this study was to model the relationship between neck electromyography (EMG) and three-dimensional (3-D) head kinematics during gaze behavior. In two monkeys, we recorded 3-D gaze, head orientation, and bilateral EMG activity in the sternocleidomastoid, splenius capitis, complexus, biventer cervicis, rectus capitis posterior major, and occipital capitis inferior muscles. Head-unrestrained animals fixated and made gaze saccades between targets within a 60° × 60° grid. We performed a stepwise regression in which polynomial model terms were retained/rejected based on their tendency to increase/decrease a cross-validation-based measure of model generalizability. This revealed several results that could not have been predicted from knowledge of musculoskeletal anatomy. During head holding, EMG activity in most muscles was related to horizontal head orientation, whereas fewer muscles correlated to vertical head orientation and none to small random variations in head torsion. A fourth-order polynomial model, with horizontal head orientation as the only independent variable, generalized nearly as well as higher order models. For head movements, we added time-varying linear and nonlinear perturbations in velocity and acceleration to the previously derived static (head holding) models. The static models still explained most of the EMG variance, but the additional motion terms, which included horizontal, vertical, and torsional contributions, significantly improved the results. Several coordinate systems were used for both static and dynamic analyses, with Fick coordinates showing a marginal (nonsignificant) advantage. Thus, during gaze fixations, recruitment within the neck muscles from which we recorded contributed primarily to position-dependent horizontal orientation terms in our data set, with more complex multidimensional contributions emerging during the head movements that accompany gaze shifts. These are crucial components of the late neuromuscular transformations in a complete model of 3-D head-neck system and should help constrain the study of premotor signals for head control during gaze behaviors.     

Postjunctional characteristics of the endplates in mammalian fast and slow muscles

We have studied the postjunctional characteristics of motor endplates in the extensor digitorum longus (EDL) and soleus muscles of the rat. At voltage clamped endplates, equilibrium interactions between acetylcholine (ACh) and the ACh receptor were determined from the dose-response curves obtained by quantitative ionophoresis of ACh. These results showed that the maximum ACh induced conductance change per unit endplate surface,g _max, was 21.8±0.9 nS/μm² in EDL and 8.2±0.9 nS/μm² in soleus, the apparent dissociation constant,K, was 65.9±4.3 μM in EDL and 43.5±3.3 μM in soleus, and the Hill-coefficient,n _H, was 2.3±0.1 in EDL and 2.2±0.1 in soleus. Single channel characteristics were derived from analysis of the ACh-induced endplate current noise. The results showed that at room temperature the mean conductance of the single channel, γ, was 24.6±1.2 pS in EDL and 23.9±1.2 pS in soleus, and the mean life time of the channel, τ, was 0.80±0.05 ms in EDL and 0.71±0.03 ms in soleus. Of all the properties studied, the maximum conductance per unit endplate surface,g _max, was significantly smaller at the soleus endplate than at the EDL endplate. The calculated density of functional ACh receptors was 62% less, and the total number of the functional ACh receptors was 60% less at the soleus endplates than at the EDL endplates. These results suggest that the soleus has a lower margin of safety for neuromuscular transmission than the EDL.     

Kinematics of the freely moving head and neck in the alert cat

 In this study we examined connections between the moment-generating capacity of the neck muscles and their patterns of activation during voluntary head-tracking movements. Three cats lying prone were trained to produce sinusoidal (0.25 Hz) tracking movements of the head in the sagittal plane, and 22.5o and 45o away from the sagittal plane. Radio-opaque markers were placed in the cervical vertebrae, and intramuscular patch electrodes were implanted in five neck muscles, including biventer cervicis, complexus, splenius capitis, occipitoscapularis, and rectus capitis posterior major. Videofluoroscopic images of cervical vertebral motion and muscle electromyographic responses were simultaneously recorded. A three-dimensional biomechanical model was developed to estimate how muscle moment arms and force-generating capacities change during the head-tracking movement. Experimental results demonstrated that the head and vertebrae moved synchronously, but neither the muscle activation patterns nor vertebral movements were constant across trials. Analysis of the biomechanical model revealed that, in some cases, modification of muscle activation patterns was consistent with changes in muscle moment arms or force-generating potential. In other cases, however, changes in muscle activation patterns were observed without changes in muscle moment arms or force-generating potential. This suggests that the moment-generating potential of muscles is just one of the variables that influences which muscles the central nervous system will select to participate in a movement. Received: 27 August 1996 / Accepted: 18 December 1996     

Neck muscle synergies during stimulation and inactivation of the interstitial nucleus of Cajal (INC)

The interstitial nucleus of Cajal (INC) is thought to control torsional and vertical head posture. Unilateral microstimulation of the INC evokes torsional head rotation to positions that are maintained until stimulation offset. Unilateral INC inactivation evokes head position-holding deficits with the head tilted in the opposite direction. However, the underlying muscle synergies for these opposite behavioral effects are unknown. Here, we examined neck muscle activity in head-unrestrained monkeys before and during stimulation (50 muA, 200 ms, 300 Hz) and inactivation (injection of 0.3 mul of 0.05% muscimol) of the same INC sites. Three-dimensional eye and head movements were recorded simultaneously with electromyographic (EMG) activity in six bilateral neck muscles: sternocleidomastoid (SCM), splenius capitis (SP), rectus capitis posterior major (RCPmaj.), occipital capitis inferior (OCI), complexus (COM), and biventer cervicis (BC). INC stimulation evoked a phasic, short-latency ( approximately 5-10 ms) facilitation and later ( approximately 100-200 ms) a more tonic facilitation in the activity of ipsi-SCM, ipsi-SP, ipsi-COM, ipsi-BC, contra-RCPmaj., and contra-OCI. Unilateral INC inactivation led to an increase in the activity of contra-SCM, ipsi-SP, ipsi-RCPmaj., and ipsi-OCI and a decrease in the activity of contra-RCPmaj. and contra-OCI. Thus the influence of INC stimulation and inactivation were opposite on some muscles (i.e., contra-OCI and contra-RCPmaj.), but the comparative influences on other neck muscles were more variable. These results show that the relationship between the neck muscle responses during INC stimulation and inactivation is much more complex than the relationship between the overt behaviors.     

Spatial orientation of extraocular muscle EMG responses to tilt in the rabbit during postnatal development

Unanaesthetized pigmented rabbits 2–45 days of age were gradually tilted (15° steps) over 360° around three orthogonal body axes. The multiunit electromyogram activity of superior rectus (SR) and superior oblique (SO) extraocular muscles was recorded with chronically implanted pairs of fine wire electrodes after each tilt step. The integrated EMG activity of the extraocular muscles in response to static tilt stimulation was trigonometrically related to tilt angle in all age groups. Each eye muscle was characterized by a typical locus of tilt positions which resulted in maximum integrated EMG activity. This locus was described by a vector of tonic vestibular activation (TVA) within the head coordinate system. The orientation of these TVA vectors within the coordinate system of the rabbit's head and, therefore, the coordination of tonic eye muscle activation in response to the position of the animal in the gravitational field did not change with postnatal age in the rabbit. Therefore, eye muscles are already capable of responding to static vestibular stimulation for the stabilization of gaze when visual information becomes available to the rabbit pups. The EMG responses to static tilt stimulation were principally determined by the scalar product of the gravity vector and the muscle's TVA vector. The response of the extraocular muscles to this effective gravitational stimulus was unchanged during the postnatal development of the rabbit: excitatory stimuli resulted in an approximately linear increase of the integrated EMG activity, inhibitory stimuli resulted in a smaller decrease to a minimum value. The correlation between the effective strength of excitatory gravity stimuli and the EMG activity of the extraocular muscles increased significantly at the time of postnatal eye opening. Since the strongest responses of SR and SO to static tilt were observed in intermediate roll-pitch positions that correspond closely to the planes of the vertical semicircular canals, a common reference frame for the cooperation of maculo-ocular and ampullo-ocular reactions is suggested.     

Motor axon sprouting in frog sartorius muscles is not altered by contralateral axotomy

Summary Sartorius muscles of the frogRana pipiens were used to study the incidence of motor nerve sprouting in normal unoperated muscles, in experimental muscles contralateral to axotomy of the sartorius nerve, and in sham-operated control muscles. Muscles were stained with either a combination of nitroblue tetrazolium nerve terminal stain and cholinesterase stain or with a combination of silver nerve terminal stain and cholinesterase stain. Each endplate that could be clearly seen was classified into one or more of the following categories: normal endplates without sprouts, three types of terminal sprouts, preterminal sprouts, nodal sprouts, sprouts of unknown origin and destination, and doubly innervated gutters. A quantitative study of 318 endplates from nine unoperated muscles, 779 endplates from 45 experimental muscles, and 694 endplates from 41 control muscles showed that all muscles had a high incidence of motor nerve sprouting and other forms of remodelling (20–28% of all endplates). There were, however, no significant differences between experimental, control, and unoperated muscles when results obtained with the same stains were compared. Results obtained with the two different stains were only slightly different. We conclude that sprouting is a very common but highly variable feature of normal frog neuromuscular junctions, and in the sartorius, contralateral axotomy does not alter this ongoing remodelling.     

Functional coupling of the stabilizing eye and head reflexes during horizontal and vertical linear motion in the cat

Summary The otolith contribution and otolith-visual interaction in eye and head stabilization were investigated in alert cats submitted to sinusoidal linear accelerations in three defined directions of space: up-down (Z motion), left-right (Y motion), and forward-back (X motion). Otolith stimulation alone was performed in total darkness with stimulus frequency varying from 0.05 to 1.39 Hz at a constant half peak-to-peak amplitude of 0.145 m (corresponding acceleration range 0.0014–1.13 g) Optokinetic stimuli were provided by sinusoidally moving a pseudorandom visual pattern in the Z and Y directions, using a similar half peak-to-peak amplitude (0.145 m, i.e., 16.1°) in the 0.025–1.39 Hz frequency domain (corresponding velocity range 2.5°–141°/s). Congruent otolith-visual interaction (costimulation, CS) was produced by moving the cat in front of the earth-stationary visual pattern, while conflicting interaction was obtained by suppressing all visual motion cues during linear motion (visual stabilization method, VS, with cat and visual pattern moving together, in phase). Electromyographic (EMG) activity of antagonist neck extensor (splenius capitis) and flexor (longus capitis) muscles as well as horizontal and vertical eye movements (electrooculography, EOG) were recorded in these different experimental conditions. Results showed that otolith-neck (ONR) and otolith-ocular (OOR) responses were produced during pure otolith stimulation with relatively weak stimuli (0.036 g) in all directions tested. Both EMG and EOG response gain slightly increased, while response phase lead decreased (with respect to stimulus velocity) as stimulus frequency increased in the range 0.25–1.39 Hz. Otolith contribution to compensatory eye and neck responses increased with stimulus frequency, leading to EMG and EOG responses, which oppose the imposed displacement more and more. But the otolith system alone remained unable to produce perfect compensatory responses, even at the highest frequency tested. In contrast, optokinetic stimuli in the Z and Y directions evoked consistent and compensatory eye movement responses (OKR) in a lower frequency range (0.025–0.25 Hz). Increasing stimulus frequency induced strong gain reduction and phase lag. Oculo-neck coupling or eye-head synergy was found during optokinetic stimulation in the Z and Y directions. It was characterized by bilateral activation of neck extensors and flexors during upward and downward eye movements, respectively, and by ipsilateral activation of neck muscles during horizontal eye movements. These visually-induced neck responses seemed related to eye velocity signals. Dynamic properties of neck and eye responses were significantly improved when both inputs were combined (CS). Near perfect compensatory eye movement and neck muscle responses closely related to stimulus velocity were observed over all frequencies tested, in the three directions defined. The present study indicates that eye-head coordination processes during linear motion are mainly dependent on the visual system at low frequencies (below 0.25 Hz), with close functional coupling of OKR and eye-head synergy. The otolith system basically works at higher stimulus frequencies and triggers Synergist OOR and ONR. However, both sensorimotor subsystems combine their dynamic properties to provide better eyehead coordination in an extended frequency range and, as evidenced under VS condition, visual and otolith inputs also contribute to eye and neck responses at high and low frequency, respectively. These general laws on functional coupling of the eye and head stabilizing reflexes during linear motion are valid in the three directions tested, even though the relative weight of visual and otolith inputs may vary according to motion direction and/or kinematics.     

Differences in activation patterns in elbow flexor muscles during isometric,concentric and eccentric contractions

Summary To investigate the relative activation of the synergistic muscles during three different types of muscle contraction, the electromyograms (EMG) of two elbow flexor muscles, the biceps brachii (BB) and the brachioradialis (BR), have been compared. To accomplish this eight healthy human subjects performed the following elbow flexions against the same load — concentric, eccentric and isometric contractions. The isometric contractions were performed at three elbow angles: 10, 45 and 90° (0° equal to full expension). The EMG were recorded by bipolar surface electrodes, and the relative activation between the two muscles was evaluated as the quotient of mean EMG activities (BR/BB). For the isotonic elbow flexions, BR/BB were calculated at three angle divisions: 0–30°, 30–60° and 60–90°. Results indicated that the relative activation of the BR during the concentric contractions was higher than that of the eccentric contraction, particularly at the extended elbow angles, i.e. the BR/BB of the concentric contractions for the elbow joint angles ranging from 0–30° and 30–60° were significantly greater (P<0.05) than those of the eccentric contractions. During the isometric and eccentric contractions, the BR/BB at the flexed joint angles tended to be greater than those at the extended angles. In contrast, there were no angle-dependent BR/BB variations during the concentric elbow flexions. Further, changing patterns in the EMG power spectra due to the type of contraction were different between BB and BR. These results indicated that the activation pattern in the two elbow flexor muscles varied with the muscle contraction pattern.